Plant Growth Regulator Uses on Citrus - ACS Symposium Series (ACS

Jul 23, 2009 - Florida Department of Citrus, Lake Alfred, FL 33850. Bioregulators. Chapter 11, pp 113–126. Chapter DOI: 10.1021/bk-1984-0257.ch011. ...
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11 Plant Growth Regulator Uses on Citrus

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W.C.WILSON Florida Department of Citrus, Lake Alfred, FL 33850

Plant Growth Regulators (PGR's) are used on c i t r u s worldwide and active research programs are in progress in most citrus-growing countries. P r i n c i p a l uses f o r PGR's on c i t r u s are for control of fruit maturity, reduction of fruit drop, prevention of rootstock sprouting, fruit thinning, preservation of fresh fruit peel q u a l i t y , and control of the abscission process for harvesting. However, there is a tremendous p o t e n t i a l for PGR's f o r freeze protection (cold hardiness), improving i n t e r n a l and external fruit color and q u a l i t y , improving storage life of fruit, increasing kg solids/ha, increasing vitamin C, c o n t r o l l i n g vegetative growth and inducing flowering and fruit s e t . Because c i t r u s is grown under a wide v a r i e t y of c l i m a t i c conditions, t h i s can greatly affect the performance of a PGR chemical on a given cultivar. A tremendous amount of research has been conducted worldwide with plant growth regulators (PGR's) on v i r t u a l l y every crop. Unlike herbicides (the "cousins of PGR s), vast commercial development has not been as rapid, as the plant has been slow in y i e l d i n g i t s growth regulating secrets. I t would appear that s e l e c t i v e k i l l i n g of plants is easier than c o n t r o l l i n g t h e i r functions and a c t i v i t i e s . Nevertheless, s i g n i f i c a n t breakthroughs have been achieved on many crops, including c i t r u s , and i t appears that commercial usage of these materials will increase in the future. Many reasons have been advanced as to why PGR s have not found markets as r a p i d l y as other plant protection chemicals. My own experience using abscission agents indicates there are several reasons, including v a r i a b i l i t y in chemical a c t i v i t y ; d i f f e r i n g c l i m a t i c conditions and c u l t i v a r s ; basic costs of chemicals and applications; phytotoxicity and human t o x i c i t y problems. Citrus production is l i m i t e d primarily by freezing temperatures because the trees will grow under a wide v a r i e t y of t r o p i c a l and subtropical conditions (1). Numerous c u l t i v a r s of sweet orange (Citrus sinensis (L.) Osbeck), grapefruit (C. paradisi Macf.), 11

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mandarin (tangerine or C. r e t i c u l a t a Blanco), lemon (fJ. limon (L.) Burm. f . ) , lime (C. a u r a n t i f o l i a Swing.) and citrons (£. medica L.) are grown worldwide. PGR spray applications reported in the numerous c i t a t i o n s were usually chemically d i l u t e concentrations. Spray coverage was usually thorough (complete). Concentrate sprays can reduce fixed spraying costs for many pesticides, but our experience with PGR s is that most have reduced effectiveness i f concentrated (2, 3). Further, our experience with surfactants and wetting agents has shown that they are b e n e f i c i a l to some PGR's, but deterimental to others (2, _3 and see i n f r a ) . P r i n c i p a l Commercial Uses of PGR s on Citrus F r u i t Thinning. Most commercial c i t r u s c u l t i v a r s do not require thinning, p a r t i c u l a r l y i f the crop is destined for processing u t i l i zation. However for fresh fruit use, s i z e , color, and other features are often important s e l l i n g points. Some c u l t i v a r s , mandarin in p a r t i c u l a r , tend to alternate bearing, thus producing heavy crops of small fruit one season but l i g h t crops of excessively large fruit the following season. Therefore, thinning to increase size ( i n the "on" year) and reduce alternate bearing is often an important h o r t i c u l t u r a l consideration. The Japanese public is very conscious of fresh fruit quality and is w i l l i n g to pay a premium for i t . Therefore, thinning to improve fruit quality ( p a r t i c u l a r l y size) of satsumas is regularly practiced (4). A leaf to fruit r a t i o of 25:1 is desired, and the purpose of chemical thinning is to reduce the amount of hand t h i n ning necessary to achieve t h i s r a t i o . The most e f f e c t i v e material has been NAA (1-napthaleneacetic acid) applied 25 days following f u l l bloom. This caused about a 30% increase in fruit drop and e f f e c t i v e l y thinned the crop 05, 6). Thinning with NAA in Japan is affected by climatic conditions, p a r t i c u l a r l y temperature (4). Best results occurred when post spray temperatures were 77°F (25^C.). High humidities also increased thinning, presumably by causing increased chemical uptake. A new growth regulator, Figaron (IZAA or ethyl-5-chloro-H-3 indazolyl-acetate), is registered for use in Japan for thinning and quality improvement of satsumas 04, 5). IZAA causes no phytoxicity. With satsuma, i t is also reported to advance fruit color and to increase j u i c e Brix (sugar content). Tests with the compound in F l o r i d a , however, have been i n e f f e c t i v e (3). Ethephon (2-chloroethylphosphonic acid) has also been tested for thinning satsumas in Japan (4, 6^), but caused excessive d e f o l i a t i o n . G i b b e r e l l i c acid (GA) applied p r i o r to bloom e f f e c t i v e l y thinned satsumas by decreasing flowering (4_) . In the USA ( F l o r i d a ) , Wheaton (7) reported the p r i n c i p a l need to thin f r u i t s has been with the c u l t i v a r s 'Dancy' and 'Murcott tangerine, both of which tend to alternately bear and produce excess i v e l y heavy crops of small-sized fruit during the "on" year. NAA was the most e f f e c t i v e and consistent material tested on both c u l t i vars. The auxins CPA (3-chlorophenoxy acetic a c i d ) , 2,4-D (2,4dichlorophenoxy acetic acid) and 2,4,5-T (2,4,5-trichlorophenoxy acetic acid) were also e f f e c t i v e thinning agents, although 2,4,5-T sometimes tended to overthin and cause phytotoxicity. In tests with ethylene-releasing materials, ethephon was 1

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generally e f f e c t i v e with 'Dancy' and caused a minimum leaf drop, but 'Murcott' showed considerable s e n s i t i v i t y to i t and over-thinning was a problem (7). GA applied prebloom provided f a i r thinning of 'Dancy' but was i n e f f e c t i v e on 'Murcott'. F r u i t thinning tests in other parts of the world with mandarintype f r u i t s have been s i m i l a r to those mentioned (8, 9_, 10) . In some parts of the world, alternate bearing of the 'Valencia' orange warrants correction by fruit thinning. In A u s t r a l i a , thinning was accomplished with ethephon applied when fruit was 1.0-1.5 cm in diameter (about 4 weeks following bloom) (11, 12). Also, GA has proven to be an e f f e c t i v e means to control f r u i t i n g of 'Valencia' when the spray was applied in winter p r i o r to the "on bloom" year (12). The GA application increased the number of vegetative buds vs. flower buds, and although cropping during the "on year" was not subs t a n t i a l l y reduced in some cases, cropping during the " o f f year" was substantially increased. Hand-thinning studies have shown that removal of only 22% of the flowers in the "on" year would greatly reduce crop differences in the two-year cycle (13). The alternate bearing problem does not appear to be of major importance in areas where the bulk of the fruit is u t i l i z e d for processing purposes (3). Preharvest Drop Control and Storage of F r u i t on the Tree. In F l o r i d a , an application of 2,4-D is recommended to prevent dropping of 'Pineapple', seedlings and 'Temple' oranges, and seedless grapefruit (14). Dilute (only) sprays should be applied in November or December. In C a l i f o r n i a and other citrus-growing areas of the world, the use of 2,4-D has generally been more successful than in F l o r i d a and is reported to prevent preharvest fruit drop of most c u l t i v a r s (9, 15, 16). Because of the importance of the Japanese market to F l o r i d a , prevention of grapefruit aging and a means of lengthening the season are important considerations (17). Therefore, should i n h i b i t i o n of color change and prevention of peel aging of seedless (sparsely seeded) grapefruit be required, a combination of 2,4-D and GA can be used (14, 18). Similar results with export grapefruit have been reported from A u s t r a l i a and South A f r i c a (19, 2CJ, 21, 22). Although delayed or late season harvesting of 'Valencia' may depress y i e l d s the following year, this reduction has not been observed with seedless ('Marsh') grapefruit (23). Preservation of Fresh F r u i t Peel Quality. There are a number of peel disorders which can be a l l e v i a t e d by applications of s p e c i f i c plant growth regulators (24). I t should be pointed out, however, that problems with peel are of primary interest to growers and shippers of fresh fruit just as i n t e r n a l q u a l i t i e s are of prime concern to processors. With lemons grown in C a l i f o r n i a , and probably most other Mediterranean climate areas of the world, peak production occurs during winter and spring when demand is low. Most lemons are harvested, stored (cured), then packed and shipped to market much l a t e r . However, during the summer when fresh lemon sales are strongest, the amount of fruit available is lowest. Therefore, i t is desirable to extend the lemon harvest season by prevention of premature yellowing (senescence) of the fruit. GA applied in the f a l l is reported to be the most b e n e f i c i a l treatment to delay o v e r a l l maturity (and yellowing) (25). These treatments also affect the second-year harvest

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pattern, probably from the influence of the GA on flowering, r e s u l t ing in more trees producing fruit during summer when market demand is high (9). Benefits of the GA treatment to lemons were better f l e x i b i l i t y in harvest patterns, a longer storage life, and a reduction in the number of small, yellow lemons (9). Navel oranges in the Central Valley of C a l i f o r n i a have r i n d softening problems which develop as the fruit matures and peel color changes from green to orange (9). These senescence changes can contribute^to a number of other rind disorders such as rind staining, sticky r i n d , puffy peel, and water spot (also considered to be weather related) (24). GA treatments applied in the early f a l l lessen or prevent these conditions, thus allowing a more orderly fresh fruit marketing season. Creasing a f f e c t s many commercial c i t r u s c u l t i v a r s throughout the world. I t is the most important single cause for r e j e c t i o n of 'Valencia* fruit in I s r a e l i packinghouses, causing a 26% discard rate (26). The use of GA applied when fruit is 3-4 cm in diameter (about July, or 4 months following anthesis) caused considerable reduction in the incidence of creasing without impeding good color development. The mode of action of GA was believed to be through renewal of growth a c t i v i t y in the affected tissues. In South A f r i c a , creasing of navel orange can cause fresh fruit packinghouse losses as high as 50% near the end of the shipping season (27, 28). The recommended control measure is to apply GA when young fruit is 30—50 mm in diameter (70-100 days after anthesi»s). The peel of mandarin fruit often tends to puff. In Japan, GA applied to satsuma mandarin (2 a p p l i c a t i o n s ) , reduced the amount of puffy fruit but also slowed the rate of chlorophyll degradation (29). F r u i t Size. Increased fruit s i z e is desirable for some c u l t i v a r s as small sizes often cause serious p r o f i t losses for fresh fruit packinghouses which must eliminate them in order to market fruit of a s p e c i f i e d l e g a l s i z e . Mandarin-type f r u i t s are often thinned with growth regulators for this purpose (see supra). For fruit destined for processing, however, size is usually not a consideration as increasing pounds s o l i d s per acre (kilograms s o l i d s per hectare) is paramount. An extensive review of a l l factors a f f e c t i n g fruit s i z e and suggested methods for i t s improvement has recently been completed by G i l f i l l i a n (30) in South A f r i c a . Lemons tend to produce t h e i r heaviest crops during the winter and spring when consumer demand is low (previously mentioned). Therefore, i t is advantageous to produce fruit which can be picked and stored u n t i l the summer period when demand (and prices) are high. In C a l i f o r n i a (31), GA and/or additional potassium f e r t i l i z e r applications are used to produce the larger, greener f r u i t s which can be harvested and stored up to 6 months. The small, yellow, treeripened f r u i t s are removed during the winter-spring period and primarily u t i l i z e d for processed products. In A u s t r a l i a (32), when lemons are held on the tree into the late spring, many grow to a s i z e which is not acceptable in the marketplace. The fruit tends to be orange-yellow which is not favored by buyers. A preharvest combination application of GA and CCC applied in the f a l l delayed fruit coloring and arrested fruit growth, thus extending the harvest into late spring and furnishing a larger quantity of small-sized, premium-quality fruit (32).

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Postharvest Treatments, F r u i t Shipment and Storage. The c i t r u s fruit is f u l l y ripe at harvest, i t contains p r a c t i c a l l y no starch reserves, and i t is not a good candidate for controlled atmospheric storage (33). Although i t is suggested that the best place to store c i t r u s probably is on the tree, there are occasions when i t must be harvested and stored, or transported very long distances in the hold of ships. Most c u l t i v a r s have a long harvest period and fruit response to plant growth regulators can vary depending on the time during the season i t was harvested, the time of the day i t was harvested, tree condition, rootstock, location, year-to-year weather variations, etc. In assessing the performance of plant growth regulators on c i t r u s , some or many of these varying conditions may affect a c t i v i t y of the chemical. Ethephon has been used to substitute f o r the effects of ethylene degreening treatments for fruit. In F l o r i d a (34, 35), dipping lemons in an ethephon solution was found to hasten the development of marketable color by 5-14 days. There was no s i g n i f i c a n t e f f e c t on the amount of decay. However, degreening mandarins with ethephon usually is more successful when applied as preharvest sprays (36). In South A f r i c a , however, postharvest ethephon dips of oranges are being used commercially in l i e u of ethylene treatments (37). The problem with lime is the opposite of lemon since i t is desired to r e t a i n the green fruit color which the public associates with limes (38). In C a l i f o r n i a , as well as s i m i l a r lemon-growing areas, preservation of the calyx tissue ( b u t t o n ) on the fruit prevents stem-end rot fungus from entering through the abscission layer tissue. This can be successfully controlled by fruit dips in aqueous solutions of 2,4-D (33), or by applications of fruit waxes containing 2,4-D or 2,4,5-T (9). Preservation of the button also improves the cosmetic appearance of the fruit, and many believe this manifests i t s freshness to the housewife. A very good Japanese market for grapefruit has developed in recent years. Because of the long t r a n s i t time in the shiphold (about 3-5 weeks from F l o r i d a ) , shipping losses can, at times, be substantial. F l o r i d a grapefruit is subject to c h i l l i n g injury (33), p a r t i c u l a r l y fruit harvested in early f a l l . As previously mentioned, (14, 18, 19, 20) preharvest treatments with 2,4-D and GA can reduce peel senescence. Attempts to control c h i l l i n g injury with plant growth regulators, however, have produced c o n f l i c t i n g results (39). The presence of seeds in citrus fruit can occasionally cause problems (40, 41). Grapefruit which are held very l a t e into the shipping season often have sprouted seeds. Preharvest treatments with 2,4-D and GA, previously mentioned, reduced the number of sprouted seeds (17, 18), but recent research was unable to confirm this (42). Reduction of A c i d i t y . Improvement of grapefruit flavor has been practiced for many years in some areas through the use of arsenate (14). The most e f f e c t i v e application period seems to be 1 to 6 weeks following bloom; however, i t can be applied as late as 4 months post bloom. Arsenate causes a reduction in t o t a l a c i d i t y of grapefruit and, consequently, causes an increase in Brix/acid r a t i o . I t is also e f f e c t i v e on oranges, but i t s effect is usually so pronounced that most of the treated fruit are i n s i p i d l y sweet. Most organic f

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and inorganic arsenical compounds will cause a c i d i t y reduction in c i t r u s f r u i t s (3), but, so f a r , no nonarsenical compounds have been i d e n t i f i e d which appear to have commercial p o s s i b i l i t i e s (43). In South A f r i c a , arsenical sprays have also been tested extens i v e l y and have been used commercially for many years (44, 45, 46). South African oranges, p a r t i c u l a r l y the 'Valencia', tend to produce fruit which has high acid content; therefore, their research e f f o r t s have been aimed at reducing a c i d i t y in orange c u l t i v a r s as w e l l as grapefruit. Recent research with arsenicals in the saline waters of the Sundays River Valley has shown that the t o t a l quantity of arsenic used can be reduced by a c i d i f y i n g the tank mix to pH 4 with s u l f u r i c acid (47, 48). (Phosphoric acid (48), however, caused considerable phytotoxicity when used.) Calcium arsenate is the preferred form (46). Arsenic residues on fruit are n e g l i g i b l e 03, 48, 49), with the largest proportion found in the peel and p r a c t i c a l l y none in the pulp and j u i c e 048, 49). In the past, inorganic arsenic has been considered to be a carcinogen and EPA clearance (or maintenance of existing clearances) of arsenicals on food crops in the United States has been quite d i f f i c u l t . However, recent n u t r i t i o n a l research throughout the world has established arsenic as an e s s e n t i a l nutrient for certain animal species (50). Although arsenic has not been proven to be e s s e n t i a l for humans, future research may, indeed, confirm these findings. If so, the very low residue levels noted for arsenictreated c i t r u s fruit, instead of presenting a health hazard, might actually be shown to be b e n e f i c i a l . Control of the Abscission Process for Mechanical F r u i t Harvesting. This subject has recently been reviewed by Wilson et a l . (51). Although c i t r u s fruit can be successfully harvested without abscission chemicals, f i e l d experience has shown that chemical loosening is desirable because less tree shaking time is required, r e s u l t i n g in less physical abuse to the machinery and trees. However, chemicals have not allowed the construction of less-powecful shakers because chemically induced fruit loosening is not always uniform, which results in about 10-15% adhering strongly to the tree. Although abscission chemicals are technically c l a s s i f i e d as growth regulators, most function by causing s u p e r f i c i a l peel burn f o l lowed by the fruit producing wound ethylene. The l a t t e r moves in some manner through tissue and a f f e c t s the abscission zone. The only commercially available chemical that appears to function through absorption by tree and fruit, followed by conversion of the chemical into ethylene, is ethephon. Temperature is the most important physiological factor a f f e c t i n g abscission of early and midseason oranges (52). Prediction of abs c i s s i o n a c t i v i t y is complicated (52), but generally good abscission a c t i v i t y occurs i f d a i l y high temperatures are 65°F (18.3 C.) or greater following spray application. Daily high temperatures lower than t h i s figure usually r e s u l t in lessened abscission a c t i v i t y or none at a l l . R a i n f a l l within 24 hr of a spray application often will negate i t s e f f e c t . The 'Valencia' orange is harvested in F l o r i d a from A p r i l u n t i l early July and in C a l i f o r n i a , Spain and other Northern Hemisphere

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citrus-growing areas from A p r i l through October. I t d i f f e r s from most f r u i t s because i t s maturity occurs 13-18 months follow­ ing bloom, which means that both a mature and immature crop are usually present when the crop is harvested. Cold temperatures are rarely a l i m i t i n g factor during harvest. However, 'Valencia* under­ goes a period of lessened physiological response to abscission chemicals which lasts 2-3 weeks and usually occurs about the f i r s t of May (54). During this period the chemical is e s s e n t i a l l y useless for loosening fruit. Before t h i s period, the fruit is very responsive to abscission chemicals, but the desired Brix/acid r a t i o s ( f r u i t matu­ r i t y ) for processing oranges has not been achieved (55) . Following the period, the immature fruit on the tree usually will average 1.52.0 cm in diameter and will have achieved s u f f i c i e n t mass so that any mechanical shaking device may remove excessive quantities along with mature fruit. Five chemicals and chemical combinations are available for use in F l o r i d a (51), and a combination of 2 of them has been used successfully. A summary of their uses is as follows: Ethephon (Ethrel) is cleared by the EPA (Environmental Protec­ tion Agency of the United States government) for use in F l o r i d a on tangerines and tangerine hybrids (14). This chemical, in addition to producing fruit loosening, enhances fruit color development. Cycloheximide (Acti-Aid) is cleared by EPA for use in F l o r i d a on oranges intended for processing. This chemical has generally produced good loosening of early and midseason oranges, but should not be applied after the spring growth begins, otherwise, severe phytotoxicity can r e s u l t . Unfortunately, i t s performance on the 'Valencia' orange has been unacceptable. Cycloheximide should not be used i f freezing temperatures are l i k e l y to occur because i t reduces the cold hardiness of the tree for an undetermined period of time. 5-chloro-3-methyl-4-nitro-lH-pyrazole (Release). This compound is available in F l o r i d a for use under experimental permit on oranges destined for processing. The chemical was the f i r s t which showed the a b i l i t y to loosen mature 'Valencia' oranges while causing v i r t u a l l y no injury to bloom, young fruit or foliage when used as recommended. Glyoxal Dioxime (Pik-Off) is a chemical very similar in mode of action to Release but i t s experimental use permit has not been renewed. Chemical Combinations: Two-way combinations of Release and cycloheximide applied with surfactant as d i l u t e sprays have given better fruit loosening than either chemical used alone. In A u s t r a l i a , abscission tests have generally reported removal force reductions very similar to those noted in F l o r i d a (56, 57, 58). Their p r i n c i p a l c u l t i v a r of i n t e r e s t is the 'Valencia' because the navel orange, although grown extensively in A u s t r a l i a ( and to a much lesser extent in Florida) is processed to a very limited extent throughout the world. The tasteless precursor of limonin, namely limonic acid Α-ring lactone, is found in raw navel oranges but upon processing, limonin is formed which produces a b i t t e r taste in the products (59). The 'Valencia' orange, early and midseason oranges grown in F l o r i d a contain limonin but at r e l a t i v e l y low l e v e l s . Minor Commercial Uses and P o t e n t i a l l y Rewarding Research Areas Control of Flowering. The i n i t i a t i o n and development of flowers

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involves a large number of i n t e r r e l a t e d , well-coordinated growth, senescence and abscission processes (60). The climate in which c i t r u s grows affects i t s flowering c h a r a c t e r i s t i c s . In t r o p i c a l , moist regions where continuous flowering, fruit set and fruit development tend to take place, flower formation usually is r e l a t i v e l y l i g h t at any one time unless drought conditions intervene to stop growth. In subtropical regions, the advent of spring growing conditions normally produces a single bloom period which may be heavy or l i g h t depending on preceding climatic conditions and/or crop load. Some c u l t i v a r s of lemons, limes and citrons tend to be everbearing (continuously blooming) under a l l conditions. High temperatures reverse the flowering process and the presence of fruit can i n h i b i t flower formation (61). A number of chemical compounds promote flowering of certain c i t r u s c u l t i v a r s (61, 62, 63, 64). As of this date, however, this writer knows of no commercial practice to increase flower formation of c i t r u s by use of plant growth regulators. F r u i t Set. F r u i t set in c i t r u s appears to be controlled by the process of competitive i n h i b i t i o n (61). Except for navel, most commercial orange and grapefruit c u l t i v a r s in the USA set s u f f i c i e n t crops so that the need for a fruit set chemical is not necessary. However, some mandarins and mandarin-grapefruit hybrids (tangelos) benefit from growth regulators which function as fruit-set chemicals. The c u l t i v a r s 'Orlando , 'Minneola', 'Nova' and 'Robinson' tend to set sparsely but in F l o r i d a increased fruit set has been obtained through sprays of GA applied to trees in f u l l bloom (65). Similar results were obtained in South A f r i c a where i t is recommended that a l l bearing blocks of 'Clementine' tangerines be sprayed at f u l l bloom with GA (66). F r u i t s retained by this method tend to be somewhat smaller and more sensitive to adverse climatic conditions than cross-pollinated fruit (65). Improved fruit set with navel orange and other c u l t i v a r s from PGR applications has been reported from many parts of the world (67, 68, 69, 70). Cold Hardiness. Trees of Citrus sp., although t r o p i c a l in o r i g i n , have the a b i l i t y to become cold hardened to some extent i f subjected to low, but not freezing temperatures, in the presence of l i g h t (71). There is no i n d i c a t i o n , however, that fruit can be cold hardened; hence i t s protection is r e s t r i c t e d to some means of a r t i f i c i a l heating or the elimination of i c e - i n i t i a t i n g bacteria which is currently under investigation (72). MH (maleic hydrazide) will produce cold-hardening of c i t r u s (9_, _71, 73), but in F l o r i d a , various bad side effects (delayed regrowth with malformed leaves) have prevented grower acceptance. Under northern C a l i f o r n i a (Mediterranean climate) conditions, night temperatures are s u f f i c i e n t l y low during f a l l that growth protection chemicals are not as important as in areas such as subtropical, moist F l o r i d a and subtropical a r i d Texas and Southern C a l i f o r n i a where "broken winter" conditions e x i s t . This condition is t y p i f i e d by i r r e g u l a r periods of warm or cold conditions which can prevent cold-hardening or cause early breaking of any cold hardiness achieved. Under Japanese growing conditions (moist, marine), NAA applied in l a t e autumn reduced damage from low temperatures (74). NAA delayed bud break of sweet orange seedlings up to 177 days (75). 1

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Translocation of NAA delayed unsprayed bud growth up to 150 days, but carryover i n h i b i t i o n e f f e c t into the second growing season was minimal. Preliminary tests in F l o r i d a (3) showed that both NAA and 2,4-DP delayed the growth of young (2-3 year) citrus trees. By summer, however, there were no v i s i b l e growth differences among any of the treatments. Control of Tree Growth. In commercial c i t r u s nurseries, stimul a t i n g young tree growth would be desirable so that the tree can be brought into production more quickly. Although GA will stimulate growth, i t does not appear to accomplish t h i s objective s a t i s f a c t o r i l y under C a l i f o r n i a conditions (9). However, under some conditions in F l o r i d a , i t has given s i g n i f i c a n t growth increases (76). Attempts to use NAA to i n h i b i t rootstock sprouting in nurseries have given mixed r e s u l t s , depending on which rootstock c u l t i v a r was used (77). In F l o r i d a grove operations, mature trees of sweet orange, lemon, grapefruit and tangerine c u l t i v a r s often must be severely hand pruned or machine hedged and topped to prevent tree sizes from exceeding heights of 15-20 f t . (4.6-6.1 meters). In C a l i f o r n i a , many lemons are hand or machine topped yearly to about 10 f t (3 meters) to f a c i l i t a t e harvesting and pest control (78). Although rootstocks to control tree s i z e are being tested in several c i t r u s growing areas, as yet, none are commercially a v a i l a b l e . Hence, mechanical control mechanisms are necessary; regrowth by c i t r u s is often very rapid and substantial. Because of the desire to develop a mechanical harvesting system for lemons, considerable tree size control research has been accomplished in both F l o r i d a and C a l i f o r n i a (79, 80, 8J., 82, 83, 84). A l l of the treatments, however, appear to have caused some phytotoxicity, although s i z e control was obtained by some of the treatments. A new PGR called P333 (paclobutrazol or (2RS, 3 RS) -1- (4-chlorophenyl) 4, 4-dimethyl-2 (1, 2, 4 - t r i a z o l - l - y l ) pentan-3-ol (paclobutrazol) has controlled growth on pome f r u i t s (85). Although research with t h i s compound is being conducted on c i t r u s , results have not, as yet, been reported. Sweet orange, certain tangelos, grapefruit and c e r t a i n mandarin c u l t i v a r s could also benefit from size control with PGR's. However, t h e i r regrowth problems are generally not as severe as those of lemons. A white latex paint c a l l e d Tre-Hold is marketed for control of trunk resprouting (86), and is used extensively in F l o r i d a on young citrus trees. It contains 2% ethyl ester of oC-NAA. Color. Two recent publications reviewed color research on c i t r u s (87, 88). The major carotenoid in orange and tangerine peel is B - c i t r a u r i n . Development of color (B-citraurin) is controlled by an i n t e r r e l a t i o n s h i p between fruit ethylene concentration and temperature (89). In Florida", preharvest sprays of ethephon are recommended to improve fruit color of mandarins and their hybrids (14). An addit i o n a l purpose of the sprays is to produce fruit abscission (loosening). The fruit must have achieved minimum i n t e r n a l quality requirements and 10-20% color break before this treatment can be e f f e c t i v e (14, 55). No surfactant should be used with ethephon and

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i t should not be combined with other materials containing surfactants; otherwise, increased leaf losses will occur. Our own observations following ethephon applications to oranges have consistently shown e r r a t i c performance and, frequently, heavy leaf losses (3). Similar r e s u l t s seem to have been observed worldwide. S t i l l , i t s o v e r a l l advantages outweigh i t s disadvantages for promotion of color (and abscission) with mandarin-type f r u i t s (14). Various attempts to control the leaf drop problem caused by ethephon have been made by many researchers, though little of the information seems to have been published, probably because of negative or c o n f l i c t i n g results (3). Another approach to the color problem has been through use of compounds which cause the fruit to produce lycopene rather than B - c i t r a u r i n (87, 88). The resultant fruit, then, tends to be red and resembles a tomato more than an orange. Best known of these compounds seems to be CPTA L2-(4-chlorophenylthio)-triethylamine hydrochloride^ (90). Although, in time, the public could probably be educated to accept oranges which are more "red" than "orange," one of the p r i n c i p a l problems with these chemicals is that, under f i e l d conditions, they produce an uneven or blotchy color pattern. Our tests have not shown that these compounds increase i n t e r n a l fruit color (3). Very little has been accomplished concerning improving i n t e r n a l c i t r u s fruit color with PGR applications. Although external peel color is not important for fruit used for processing, i n t e r n a l (juice) color is very important. Oranges grown in the tropics usually have poor external and only f a i r i n t e r n a l color, but i n t e r n a l q u a l i t i e s such as j u i c e content and soluble solids may be reasonably good (1, S]_ 88). However, development of a plant growth regulator to improve i n t e r n a l color, or any of these other q u a l i t i e s , would be b e n e f i c i a l . Nursery Aids The cost of a c i t r u s tree can be a substantial part of the i n i t i a l cost of planting a c i t r u s grove (orchard), hence i t would be b e n e f i c i a l to decrease the time of germination of seed, increase the per cent germination and increase the growth rate of the seedl i n g (61) . Several methods have been suggested (91, 92^, 93^ 94, 95, 96, 97). Plant growth regulators are also reported to aid propagation. The highest number of roots was obtained" from a i r layers of seedless lemon when the stem was treated with an aqueous solution containing a combination of IBA and NAA (98). Cuttings of s o f t wood or semihard wood are reported to respond well to growth regulators such as IBA (61). Attempts to increase the s i z e , p a r t i c u l a r l y g i r t h , of a seedling using growth regulators have given mixed r e s u l t s . GA applications generally tend to increase the stem length but at the expense of reduced g i r t h (99) . However, weekly applications of GA with several spreader-sticker and antitranspirant products s i g n i f i c a n t l y increased plant height, intermode length, and stem diameter of sour orange (C. aurantium L.) seedlings (76). Plant growth regulators may be useful for c o n t r o l l i n g shape of young c i t r u s trees (100, 101). They can also reduce the time lapse between bud grafting and outgrowth of the bud (9, 102, 103). 9

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Conclusion A great deal of PGR research has been conducted on c i t r u s , and some compounds have achieved commercial importance. Although PGR s continue to have problems r e l a t i n g to t h e i r application and use, most appear to be t h e o r e t i c a l l y solvable through continued research programs now in existence. For a more in depth treatment of PGR uses on c i t r u s , see Wilson (3). T

Literature Cited

Bioregulators Downloaded from pubs.acs.org by MONASH UNIV on 02/26/16. For personal use only.

1.

2. 3.

4. 5. 6. 7. 8. 9.

10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24.

Reuther, W. In "Citrus Industry"; Reuther, W. (Editor); Univ. of C a l i f . , Div. Agric. Services, Berkeley, 1973; V o l . 3, p. 280. Wilson, W. C.; Holm, R. E.; Clark, R. K. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 2, 404. Wilson, W. C. The Use of Exogenous Plant Growth Regulators on C i t r u s . In "Plant Growth Regulating Chemicals"; N i c k e l l , L. G. (Editor); CRC Press, Inc., Boca Raton, F l o r i d a , 1983; V o l . 1, Chap. 8. Iwahori, S. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 263. Hirose, K.; Iwagaki, L.; Suzuki, K. Proc. Int. Soc. Citri­ culture, 1978, 270. Iwahori, S.; Oohata, J. T. S c i . Hortic., 1976, 4, 167. Wheaton, T. A. Proc. Int. Soc. C i t r i c u l t u r e , 1981, 1, 263. Gallasch, P. T. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 276. Coggins, C. W., J r . ; Hield, H. Z. In "The Citrus Industry"; Revised ed., Reuther, W.; Webber, H. J.; Batchelor, L. D. (Editors); Univ. of C a l i f . , Div. Agric. Services, Berkeley, 1968, V o l . 1, p. 371. Monselise, S. P. S c i . Hortic., 1979, 11, 151. Gallasch, P. T.; Bevington, Κ. B.; Hocking, D.; Moss, G. I. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 273. Moss, G. I.; Bevington, Κ. B. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 260. Moss, G. Rural Research, 1977, 94, 22. Knapp, J. L. F l a . Coop. Ext. Ser., C i r c u l a r 393-1 (revised annually), 1983. Stewart, W. S.; Hield, H. Z. Proc. Am. Soc. Hortic. S c i . , 1950, 55, 163. Sarooshi, R. Α.; Stannard, M. C. Aust. J. Exp. Agric. Anim. Husb., 1975, 15, 429. Krezdorn, A. H. C a l i f . Citrogr., 1977, 62, 85. Dinar, Η. Μ. Α.; Krezdorn, A. H.; Rose, A. J. Proc. F l a . State Hortic. Soc., 1976, 89, 4. El-Zeftawi, Β. M. J. Hortic. S c i , 1980, 55, 211. G i l f i l l a n , I. M.; Koekemoer, W.; Stevenson, J. The Citrus Grower and Sub-Tropical F r u i t Journal, 1972, 458, 6. G i l f i l l a n , I. M.; Koekemoer, W.; Stevenson, J. Proc. Int. Soc. C i t r i c u l t u r e , 1973, 3, 335. G i l f i l l a n , I. M.; Stevenson, J. A. Citrus Sub-Trop. F r u i t J., 1976, 507, 5. Ramirez, J. M.; Krezdorn, A. H.; Rose, A. J. Proc. F l a . State Hortic. Soc., 1977, 90, 61. Grierson, W. Proc. Int. Soc C i t r i c u l t u r e , 1981, 2, 764.

124 25. 26. 27. 28. 29.

Bioregulators Downloaded from pubs.acs.org by MONASH UNIV on 02/26/16. For personal use only.

30.

31. 32. 33. 34. 35. 36. 37. 38.

39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55.

56.

BIOREGULATORS: CHEMISTRY AND USES Monselise, S. P. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 250. Monselise, S. P.; Weiser, M.; Shafir, N.; Goren, R.; Goldschmidt, Ε. E. J. H o r t i c . S c i . , 1976, 51, 341. Gilfillan, I. M.; Stevenson, J. Α.; Wahl, J. P. Proc. Int. Soc. C i t r i c u l t u r e , 1981, 1, 224. Gilfillan, I. M.; Stevenson, J. Α.; Holmden, E.; F e r r e i r a , C. J.; Lee, A. The Citrus and Subtrop. F r u i t J., 1980, 605, 11. Kuraoka, T.; Iwasaki, K.; I s h i i , T. J. Am. Soc. Hortic. S c i . 1977, 102, 651. Gilfillan, I. M. "Report on a visit to Japan, A u s t r a l i a , and the United States of America with Emphasis on F r u i t Size Improvement," Outspan Citrus Center, Nelspruit, South A f r i c a , 1983. Burns, R. Μ., 1982, (personal communication). El-Zeftawi, Β. M. S c i . Hortic., 1980, 12, 177. Grierson, W.; Hatton, T. T. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 1, 227. Wardowski, W. F.; Barmore, C. R.; Smith, T. S.; DuBois, C. W. Proc. F l a . State Hortic. Soc., 1974, 87, 216. Jahn, O. L. Proc. F l a . State Hortic. Soc., 1974, 87, 218. Brown, G. E.; Barmore, C. R. Proc. F l a . State Hortic. Soc., 1978, 89, 198. Gilfillan, I. Μ., 1982 (personal communication). Bleinroth, E. W.; Hansen, Η. Α.; F e r r e i r a , V. L. P.; Angelucci, E. Coletanea do Instituto de Technologia de Alimentos, 1976, 7, 343. Ismail, Μ. Α.; Grierson, W. HortScience, 1977, 12, 118. Feinstein, B.; Monselise, S. P.; Goren, R. HortScience, 1975, 10, 385. Lewin, I. J.; Monselise, S. P. S c i . Hortic., 1976, 4, 229. Ferguson, L.; Davies, F. S.; Ismail, Μ. Α.; Wheaton, T. A. Plant Growth Reg. Soc. of Am., 1983, 10, 175. Wilson, W. C.; Kenny, D. S.; Holm, R. E. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 2, 692. Crous, P. Α., Citrus Grower, 1941, 94, 1. Mynhardt, C. B. Citrus Grower, 1956, 273, 1. Basson, W. J. Farming in South A f r i c a , 1959, July, p. 52. Gilfillan, I. M.; Stevenson, J. A. The Citrus & Subtrop. F r u i t J., 1976, March, p. 13. Gilfillan, I. M.; Wahl, P.; Holmden, E.; Reay, N.; Stevenson, J. A. The Citrus & Subtrop. F r u i t J., 1975, March, p. 5. Kesterson, J. W.; Braddock, R. J.; Koo, R. C. J.; Reese, R. L., HortScience, 1975, 10, 65. Metz, W. Science, 1981, 213, 1332. Wilson, W. C.; Coppock, G. E.; Attaway, J. A. Proc. Int. Soc. C i t r i c u l t u r e , 1981, 1, 278. Biggs, R. H.; Kossuth, S. V. Acta Hortic., 1981, 120, 71. Wilson, W. C. Proc. Plant Growth Reg. Soc. of Am., 1980, 7, 41. Wheaton, Τ. Α.; Wilson, W. C.; Holm, R. E. J. Am. Soc. Hortic. S c i . , 1977, 102, 580. Wardowski, W.; Soule, J.; Grierson, W.; Westbrook, G. F l a . Cooperative Extension Services, Univ. of F l a . , B u l l . 188, 1979. Hutton, R. J. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 257.

11. 57. 58. 59. 60. 61. 62.

Bioregulators Downloaded from pubs.acs.org by MONASH UNIV on 02/26/16. For personal use only.

63. 64. 65. 66. 67. 68. 69. 70. 71. 72.

73. 74.

75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88.

89.

WILSON

Plant Growth Regulator Uses on Citrus

125

Hutton, R. J. Proc. Int. Soc. C i t r i c u l t u r e , 1981, 1, 281. El-Zeftawi, Β. M. Proc. Int. Soc. C i t r i c u l t u r e , 1978, 255. Ting, S. V.; Fisher, J. F.; Nagy, S. , 1982, (personal communication). Goldschmidt, Ε. E. Plant & Cell Physiol., 1980, 21, 193. Moss, G. I. Ciba-Geigy Agrochemicals Tech. Monograph, 1975, No. 4, p. 61. N i r , I.; Goren, R.; Leshem, B. J. Am. Soc. Hortic. S c i . , 1972, 97, 695. Goren, R.; Monselise, S. P. Planta (Berl.), 1970, 88, 364. Salomon, E. Acta Hortic., 1980, 114, 11. Krezdorn, A. H.; Jernberg, D. C. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 2, 660. de Lange, J. H.; du P l e s s i s , S. F.; Vincent, A. P.; du Preez, M. B.; Holmden, Ε. Α.; Rabe, E. Subtropica, 1982, 3, 7. Dayuan, Wang. HortScience, 1981, 16, 657. Shawki, I.; El-Tomi, Α.; Nasr, A. Egypt. J. Hortic., 1978, 5, 115. Kumar, R.; Singh, J. P.; Gupta, O. P. Haryana J. Hortic. Soc., 1975, 4, 123. Garcia-Martinex, J. L.; Garcia-Papi, M. A. Scientia Hortic., 1979, 10, 285. Yelenosky, G. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 1, 199. Lindow, S. E.; Arny, P. C.; Upper, C. D.; Barchet, W. R. In "Plant Cold Hardiness and Freezing Stress," Li, P. H.; Sakai, A. (Editors), Academic Press, New York, 1978, p. 249. Hendershott, C. H. Proc. Am. Soc. Hortic. S c i , 1962, 80, 241. Konakahara, M. Special B u l l e t i n of the Shizuoka Prefectural Citrus Experiment Station No. 3, Komagoe, Shimizu-City, Japan, 1975, 1. Nauer, E. M.; Boswell, S. B.; Holmes, R. C. HortScience, 1979, 14, 525. M i l l e r , I. Α.; Young, M. J. HortScience, 1982, 17, 673. Nauer, Ε. M.; Boswell, S. B. HortScience, 1978, 13, 166. Burns, R. Μ., 1983, (personal communication). Lundberg, E. C.; Smith, T. S. Proc. F l a . State Hortic. Soc., 1974, 87, 20. Moye, Η. Α.; Willson, A. E. Proc. F l a . State Hortic. Soc., 1977, 90, 272. P h i l l i p s , R. L.; Tucker, D. P. H. HortScience, 1974, 9, 199. Boswell, S. B.; McCarty, C. D.; Ede, L. L.; Chesson, J. H. Citrograph, 1975, 60, 405. Boswell, S. B.; Burns, R. M.; Hield, Η. Z. HortScience, 1976, 11, 115. Boswell, S. B.; Embleton, T. W.; Jones, W. W.; Summers, L. L. HortScience, 1981, 16, 41. Williams, M. W. Plant Growth Reg. Soc. of Am., 1982, 9, 78. Plant Growth Regulator Handbook, 2nd E d i t i o n , Plant Growth Reg. Soc. of Am., Lake A l f r e d , Fl, 1981, p. 78. Stewart, I. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 1, 308. Stewart, I. In "Citrus N u t r i t i o n and Quality," Nagy, S.; Attaway, J. A. (Editors); ACS Symposium Series 143; American Chemical Society, 1980, p. 129. Wheaton, T. A; Stewart, I. J. Am. Soc. Hortic. S c i . , 1973,98,337.

126

90. 91. 92. 93. 94.

Bioregulators Downloaded from pubs.acs.org by MONASH UNIV on 02/26/16. For personal use only.

95.

96. 97. 98. 99.

100.

101. 102. 103.

BIOREGULATORS: C H E M I S T R Y A N D USES

Yokoyama, H.; Hsu, W.; Poling, S. M.; Hayman, E.; De Benedict, C. Proc. Int. Soc. C i t r i c u l t u r e , 1977, 3, 717. Burns, R. M.; Coggins, C. W. C a l i f . Agric., 1969, 23, 18. Choudhari, B. K.; Chakrawar, V. R. J. Maharashtra Agric. U n i v e r s i t i e s , 1980, 5, 173. Choudhari, Β. K.; Chakrawar, V. R. J. Maharashtra Agric. U n i v e r s i t i e s , 1981, 6, 19. Shukla, K. S.; Misra, R. L.; Kaul, M. K.; Prasad, A. Haryana J. Hortic. S c i . , 1978, 7, 162. Abdalla, K. M.; Wakeel, A. T. El; Masiry, H. H. El. Research B u l l e t i n , A i n Shams Univ., Faculty of Agric. Zagazig Univ., Cairo, Egypt, 1978, No. 944. Misra, R. S.; Verma, V. K. Progressive Hortic., 1980, 12, 79. Singh, H. K.; Shankar, G.; Makhija, M. Haryana J. H o r t i c . S c i . , 1979, 8, 194. P a t i l , S. B.; Chakrawar, V. R. Punjab Hortic. J., 1979, 19, 110. Abdalla, K. M.; El-Wakeel, A. T.; Ε.-Masiry, Η. Η. Research B u l l e t i n , Faculty of Agric., A i n Shams Univ., Zagazig Univ., Zagazig, Egypt, 1979, No. 937. Abdalla, Κ. M.; Wakeel, A. T. El; Masiry, H. H. El. Research B u l l e t i n , Ain Shams Univ., Zagazig Univ., Cairo, Egypt, 1978, No. 936. Hassaballa, I. A. Research B u l l e t i n , Ain Shams Univ., Faculty of Agric., Zagazig Univ., Moshtohor, Egypt, 1979, No. 1076. M a i t i , R. B.; Singh, S. M.; Singh, I. J. Indian J. Hortic., 1959, 16, 249. Mauer, Ε. M.; Boswell, S. B.; Holmes, R. C. HortScience, 1979, 14, 229.

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